The cardiac muscle is a type of involuntary striated muscle found in the walls of the heart. As it contracts, it propels blood into the heart and through the blood vessels of the circulatory system.

Metabolism

Cardiac muscle is adapted to be highly resistant to fatigue: it has a large number of mitochondria, enabling continuous aerobic respiration via oxidative phosphorylation, numerous myoglobins (oxygen-storing pigment) and a good blood supply, which provides nutrients and oxygen. The heart is so tuned to aerobic metabolism that it is unable to pump sufficiently in ischaemic conditions. At basal metabolic rates, about 1% of energy is derived from anaerobic metabolism. This can increase to 10% under moderately hypoxic conditions, but, under more severe hypoxic conditions, not enough energy can be liberated by lactate production to sustain ventricular contractions.

Under basal aerobic conditions, 60% of energy comes from fat (free fatty acids and triacylglycerols/triglycerides), 35% from carbohydrates, and 5% from amino acids and ketone bodies. However, these proportions vary widely according to nutritional state. For example, during starvation, lactate can be recycled by the heart. This is very energy efficient, because one NAD⁺ is reduced to NADH and H⁺ (equal to 2.5 or 3 ATP) when lactate is oxidized to pyruvate, which can then be burned aerobically in the TCA cycle, liberating much more energy (ca 14 ATP per cycle).

In the condition of diabetes, more fat and less carbohydrate is used due to the reduced induction of GLUT4 glucose transporters to the cell surfaces. However, contraction itself plays a part in bringing GLUT4 transporters to the surface. This is true of skeletal muscle, but relevant in particular to cardiac muscle, since it is always contracting.

Unlike skeletal muscle, which contracts in response to nerve stimulation, specialized pacemaker cells at the entrance of the right atrium termed the sinoatrial node display the phenomenon of automaticity and are myogenic, meaning that they are self-excitable without a requisite electrical impulse coming from the central nervous system. However, the sinoatrial node is modulated by the autonomic nervous system. The rest of the myocardium conducts these action potentials by way of electrical synapses called gap junctions. It is because of this automaticity that an individual's heart does not stop when a neuromuscular blocker (such as succinylcholine or rocuronium) is administered, such as during general anesthesia.

A single cardiac muscle cell, if left without input, will contract rhythmically at a steady rate; if two cardiac muscle cells are in contact, whichever one contracts first will stimulate the other to contract, and so on. This inherent contractile activity is heavily regulated by the autonomic nervous system. If synchronization of cardiac muscle contraction is disrupted for some reason (for example, in a heart attack), uncoordinated contraction known as fibrillation can result.

Intercalated disc

An intercalated disc is an undulating double membrane separating adjacent cells in cardiac muscle fibers. Intercalated discs support synchronized contraction of cardiac tissue. They can easily be visualized by a longitudinal section of the tissue.

Three types of membrane junctions exist within an intercalated disc: fascia adherens, macula adherens, and gap junctions. Fascia adherens are anchoring sites for actin, and connects to the closest sarcomere. Macula adherens stop separation during contraction by binding intermediate filaments joining the cells together, also called a desmosome. Gap junctions allow action potentials to spread between cardiac cells by permitting the passage of ions between cells, producing depolarization of the heart muscle. When observing cardiac tissue through a microscope, intercalated discs are an identifying feature of cardiac muscle

Rate

Specialized pacemaker cells in the sinoatrial node normally determine the overall rate of contractions, with an average resting pulse of 72 beats per minute. However, in the absence of any nervous system or hormonal input, the conducting cells of the sinoatrial node exhibit an intrinsic firing rate of 100 beats per minute. This indicates that the parasympathetic division of the autonomic nervous system predominates at rest.

The central nervous system does not directly create the impulses to contract the heart, but only sends signals to speed up or slow down the heart rate through the autonomic nervous system using two opposing kinds of modulation:

• (1) sympathetic nervous system (fight or flight response)
• (2) parasympathetic nervous system (rest and repose)

Since cardiac muscle is myogenic, the pacemaker serves only to modulate and coordinate contractions. The cardiac muscle cells would still fire in the absence of a functioning SA node pacemaker, albeit in a much less efficient manner. When another part of the conducting system takes over when there is damage to the SA node, it is then referred to as an ectopic pacemaker. Note that the heart can still beat properly even if its connections to the central nervous system are completely severed.

Role of calcium

In contrast to skeletal muscle, cardiac muscle requires both extracellular calcium and sodium ions for contraction to occur. In this sense, it is intermediate between smooth muscle, which has a poorly developed sarcoplasmic reticulum and derives its calcium predominately across the sarcolemma, and skeletal muscle, which is activated by calcium stored in the sarcoplasmic reticulum (SR).

The reason for the calcium dependence is due to the mechanism of calcium-induced calcium release (CICR) from the SR that must occur under normal excitation-contraction (EC) coupling to cause contraction.

Appearance

Striation

Cardiac muscle exhibits cross striations formed by alternating segments of thick and thin protein filaments, which are anchored by segments called T-lines. In contrast to skeletal muscle, cardiac muscle cells are branched instead of linear.

Like skeletal muscle, the primary structural proteins of cardiac muscle are actin and myosin. The actin filaments are thin causing the lighter appearance of the I bands in muscle, while myosin is thicker lending a darker appearance to the alternating A bands as observed by a light enhanced microscope.

T-Tubules

Another histological difference between cardiac muscle and skeletal muscle is that the T-tubules in cardiac muscle are larger, broader and run along the Z-Discs. There are fewer T-tubules in comparison with Skeletal muscle. Additionally, cardiac muscle forms dyads instead of the triads formed between the T-tubules and the sarcoplasmic reticulum in skeletal muscle.

Intercalated Discs

Under light microscopy, intercalated discs appear as thin, typically dark-staining lines dividing adjacent cardiac muscle cells. The intercalated discs run perpendicular to the direction of muscle fibers. Under electron microscopy, an intercalated disc's path appears more complex. At low magnification, this may appear as a convoluted electron dense structure overlying the location of the obscured Z-line. At high magnification, the intercalated disc's path appears even more convoluted, with both longitudinal and transverse areas appearing in longitudinal section. Gap junctions (or nexus junctions) fascia adherens (resembling the zonula adherens), and desmosomes are visible. In transverse section, the intercalated disk's appearance is labyrinthine and may include isolated interdigitations.

References

• Indiana State University, Muscle action

See also

• Myocardium
• Heart
• Circulatory system
• Cardiac action potential
• Calcium sparks
• Troponin